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Home , Bronze Age, Land of Punt, Levant, Southern Levant, Tel Erani, Terqa

Exotic foods reveal contact between South and the Near during the second millennium BCE

Ashley Scotta,b, Robert C. Powerb,c, Victoria Altmann-Wendlingb,d, Michal Artzye, Mario A. S. Martinf, Stefanie Eisenmanna,b, Richard Hagana,b, Domingo C. Salazar-Garcíag,h, Yossi Salmone, Dmitry Yegorovi, Ianir Milevskii, Finkelsteinf, Philipp W. Stockhammera,b,1, and Christina Warinnera,j,1

aDepartment of Archaeogenetics, Max Planck Institute for the Science of , 07743 Jena, ; bInstitute for Pre- and Protohistoric and Archaeology of the Roman Provinces, Ludwig Maximilian University Munich, 80539 Munich, Germany; cDepartment of Human Evolution, Max Planck Institute for Evolutionary Anthropology, 04103 Leipzig, Germany; dInstitute for Egyptology, Julius-Maximilians-University Würzburg, 97070 Würzburg, Germany; eLeon Recanati Institute for Maritime Studies, University of , 3498838 Haifa, Israel; fInstitute of Archaeology, University, 69978 Tel Aviv, Israel; gIkerbasque-Basque Foundation for Science, Grupo de Investigación en Prehistoria IT-1223-19 (Universidad del País Vasco/Euskal Herriko Unibertsitatea), 01006 Vitoria-Gasteiz, Spain; hDepartament de Prehistòria, Arqueologia i Història Antiga, University of València, 46010 València, Spain; iArchaeological Research Department, Archaeological Division, Israel Antiquities Authority, 91004, Israel; and jDepartment of Anthropology, Harvard University, Cambridge, MA 02138

Edited by Dolores R. Piperno, Smithsonian Institution, Washington, DC, and approved November 10, 2020 (received for review July 21, 2020) Although the key role of long-distance in the transformation intensity of exchange gained momentum during the Middle and of cuisines worldwide has been well-documented since at least the Late Ages of the second millennium BCE (15–17). Roman era, the of the Eurasian food trade is less visible. During this period, bronze was produced on a large scale across In order to shed light on the transformation of Eastern Mediterra- (4, 5, 18), and urban societies and early states linked by nean cuisines during the and Early Age, we ana- these trade routes developed a rapidly growing interest in exotic lyzed microremains and proteins preserved in the dental calculus goods, including plant and animal products. In the early first of individuals who lived during the second millennium BCE in the millennium BCE, such trade networks had effectively linked Southern . Our results provide clear evidence for the con- West and , and several economically important crops sumption of expected staple foods, such as (Triticeae), ses- had become widely dispersed throughout the (Fig. 1, ame (Sesamum), and dates (Phoenix). We additionally report and SI Appendix, Fig. S1). evidence for the consumption of soybean (Glycine), probable ba- ANTHROPOLOGY nana (Musa), and turmeric (Curcuma), which pushes back the ear- Evidence for Long-Distance Trade of Material Goods and Animals. liest evidence of these foods in the Mediterranean by centuries During the second millennium BCE, textual evidence in the (turmeric) or even millennia (soybean). We find that, from the attests to a large amount of goods being transported early second millennium onwards, at least some people in the over great distances. For example, tablets from the had access to food from distant locations, Assyrian trade post of Kaneš (Kanesh) in record car- including , and such goods were likely consumed as oils, avans of hundreds of donkeys regularly transporting goods be- dried fruits, and spices. These insights force us to rethink the com- tween the Mesopotamian city of Aššur () and Central plexity and intensity of Indo-Mediterranean trade during the Anatolia during the 19th and18th centuries BCE (19, 20). Bronze Age as well as the degree of globalization in early Eastern Starting with the expansion policy of Thutmosis III ( . BCE) into the Levant, the flow of goods and people in West Asia

proteomics | Bronze Age | Eastern Mediterranean | | Significance early globalization Here we report the identification of staple and exotic food ong-distance trade across Eurasia has played a major role in remains in Bronze and Early dental calculus from the Lconnecting distant societies throughout , with . The analysis of dietary plant microremains the silk and spice trade being emblematic for early globalization and proteins sheds new light on consumed exotic foods from (1–3). Recent decades of archaeological research have demon- South and East Asia during the second millennium BCE. We strated the deep prehistory of these trans-Eurasian exchange provide the earliest direct evidence in the Mediterranean to networks and support a much earlier onset of globalization, date for the consumption of sesame, soybean, probable ba- known as “Bronzization” (4), which traces its roots to the Bronze nana, and turmeric. The recovery and identification of diverse Age during the third millennium BCE. The increasing impor- foodstuffs using molecular and microscopic techniques enables tance of bronze served as a major impetus for the establishment a new understanding of the complexity of early trade routes of extensive trade contacts (5), which were largely driven by the and nascent globalization in the and raises uneven distribution of highly valued raw materials, such as , questions about the long-term maintenance and continuity of , and (6–8). In addition to these raw mate- this trade system into later periods. rials, finished objects—as well as technologies, practices, and knowledge—were also conveyed over unprecedented distances. Author contributions: A.S., R.C.P., M.A., Y.S., P.W.S., and C.W. designed research; A.S., R.C.P., S.E., R.H., and D.C.S.-G. performed research; A.S., R.C.P., V.A.-W., M.A.S.M., D.Y., The major corridors of exchange connected Eurasia both by land I.M., I.F., P.W.S., and C.W. analyzed data; and A.S., R.C.P., V.A.-W., P.W.S., and C.W. wrote across the Eurasian steppes and and by sea from the paper. to the Near East via both the and the The authors declare no competing interest. (9, 10). These networks served to link the major Bronze Age This article is a PNAS Direct Submission. river societies of , , the Indus Valley, Published under the PNAS license. and Central to neighboring cultures in the Levant, the 1To whom correspondence may be addressed. Email: [email protected] or , the Iranian Plateau, and the Central Asian [email protected]. oases (11–13). Such trade networks also further extended into This article contains supporting information online at https://www.pnas.org/lookup/suppl/ Anatolia, the Aegean, and throughout South and East Asia. doi:10.1073/pnas.2014956117/-/DCSupplemental. Despite periodic disruptions of some trade routes (14), the Published December 21, 2020.

PNAS 2021 Vol. 118 No. 2 e2014956117 https://doi.org/10.1073/pnas.2014956117 | 1of10 Downloaded by guest on October 1, 2021 Fig. 1. Map of representative archaeobotanical evidence for the spread and trade of food crops prior to 500 BCE. See SI Appendix for data sources. Map Inset shows the location of the sites of Megiddo and on the southern Levantine coast; new dietary finds reported in this study are indicated for each site.

intensified, which is documented most prominently in the fa- Kingdom (2050 to 1710 BCE) Egyptian tomb at the site of - mous letters (21). Ancient literary sources and illus- Gebelein points in the same direction (42). At the same time, trations also reference long-distance journeys to foreign lands to other animals, such as domestic chicken (Gallus gallus - obtain exotic goods, such as ivory, ostrich eggshells, , and ticus) were brought from East Asia to the Near East (possibly via . Among the most well-known of these accounts is South Asia) and displayed as exotic curiosities and used for an expedition initiated by queen to the cockfighting. Sporadic evidence of chicken occurs in Anatolia, land of Punt (probably located in the Horn of ) in , , and Egypt as early as the third millennium BCE, and the 15th century BCE (22, 23). In addition, evidence for long- by the late first millennium BCE some sites in the Levant appear distance trade between the Near East and the Indian subconti- to have economically specialized in chicken husbandry (43). nent is growing for the second and third millennium BCE (8, 24), and includes written accounts (25–27), seals and stone weights Evidence for Long-Distance Trade of Culinary and Economic Plants. (8, 28–30), shell, lapis lazuli, and carnelian jewelry (8, 24, 31, 32), Although evidence for the movement of durable goods and an- and timber and ivory (33, 34). imals is richly attested by historical and archaeological evidence, Live animals were also transported long distances. Zooarch- direct evidence for culinary and economic plants is more limited. aeological and isotopic analyses have identified the movement of Plant remains are highly perishable, and only certain conditions donkeys from Egypt to the Southern Levant during the third lead to macrobotanical preservation, which may be biased with millennium BCE (35), and ancient DNA analysis has docu- respect to plant type or part (44). Nevertheless, plant remains, mented the transport of pigs from Italy to the southern Greek such as charred seeds, have documented the eastward spread of mainland (36) and from the Aegean to the Southern Levant wheat and barley across Eurasia during and after the late fourth during the second millennium BCE (37). In addition, depictions millennium BCE (45), as well as the westward spread of of zebu (Bos indicus), a humped subspecies of native cultivation from East Asia during the third millennium BCE (9, to South Asia (38), have been found in Mesopotamia as early as 46, 47). Macrobotanical and microbotanical evidence also con- the third millennium BCE (39), and a clear depiction of a zebu firm the establishment of citron ( medica), a fruit tree of (Fig. 2) appears on a bichrome ceramic vessel from the southern South/Southeast Asian origin (48, 49), as an important crop in Levant dating to BCE, the same period when de- the Levant and Egypt by the first millennium BCE (50–52), al- pictions of zebu also become common in Egypt (39). Zooarch- though it may have been first introduced into the Eastern aeological study of faunal remains and genetic data further Mediterranean as early as the fourth millennium BCE (53, 54). confirm the presence of zebu cattle in the Levant during the (Cucumis melo), another important crop of South Asian second millennium BCE (37, 40), and genetic evidence suggests origin (55), was also cultivated in the Near East during the possible taurine–zebu hybridization at the site of Megiddo at Bronze Age. Textual references to appear in third mil- approximately 900 BCE (37). Monkeys depicted in 18th and 17th lennium BCE Sumerian texts, and melons and cucurbits are century BCE frescoes at Akrotiri on the Aegean island of Thera depicted in Egyptian tombs from the Old Kingdom onwards (54, were recently identified as South Asian gray langurs (Semnopi- 56, 57). Cucurbit seeds have been reported in Near Eastern ar- thecus sp.), with a probable origin in the Indus valley (41), and chaeological contexts as early as the sixth century BCE (54), but the identification of dermestid khapra beetles (Trogoderma finds prior to the first millennium BCE are less secure in their granarium) that had infested a wheat deposit from a Middle taxonomic assignment (56).

2of10 | PNAS Scott et al. https://doi.org/10.1073/pnas.2014956117 Exotic foods reveal contact between South Asia and the Near East during the second millennium BCE Downloaded by guest on October 1, 2021 ANTHROPOLOGY Fig. 2. Iconographic representation of a zebu on a bichrome ceramic vessel from Tel Gerisa, Israel, 16th century BCE (photo courtesy of Zeev Herzog).

Beyond grains and fruits, there is also growing evidence for a During the Middle and Late Bronze Age, Megiddo was a spice trade between South Asia and the Eastern Mediterranean. major urban center in the Southern Levant, and it was embedded This is supported by recent findings from organic residue anal- within long-distance networks and ruled by local kings. We se- ysis, including evidence for vanillin (58) and possibly also cin- lected Megiddo because it had already yielded suggestive evi- namon (59), nutmeg, and jasmine (60), although evidence for the dence for exchange with South Asia in the form of both zebu latter three spices requires further confirmation. Whereas many genetic evidence (37) and vanillin residues (58). In this study, aspects of this early trade remain unknown, some extraordinary we analyze individuals from a variety of mortuary contexts, finds leave no doubt that an Indo-Mediterranean spice trade including pit burials, brick-lined pit burials, burials, already existed during the Bronze Age (6, 9, 59). Peppercorns masonry-constructed collective tombs, and a recently excavated used in the mummification of Ramses II in 1213 BCE, for ex- Middle Bronze Age royal tomb containing the so-called “king” ample, are native to southern India (61–63), and cloves, origi- (MGD001) and “queen” (MGD002) of Megiddo (SI Appendix). nally from Indonesia, were found at Terqa, Syria dating to 1720 Dating to ∼500 y later in time, the Tel Erani cemetery is one BCE (64, 65), having likely followed an indirect route to Mes- of the few Early Iron Age cemeteries excavated to date in Israel opotamia via South Asian trade routes (30, 66). Both of these (SI Appendix). Although the related settlement is less under- examples highlight the extent of trade during this stood, the cemetery has been associated with the “Philistine” period, despite the declining influence of the Indus Valley and the occupation at the Southern Levant from the 12th century BCE restructuring of political networks throughout the region (33). onwards. In contrast to Megiddo, where several individuals sampled by us derived from high- or highest-status burial con- Emerging Picture of a Dynamic and Complex Exchange Network. texts, the individuals from Tel Erani instead appear to represent While the details of Bronze Age trade remain patchy, the the rural general population of the Early Iron Age in this region. overall evidence points toward the existence of a dynamic and By analyzing the two sites, we aim to gain a broader perspective complex exchange network connecting the Mediterranean with on Levantine cuisines during the second millennium BCE, a pe- South Asia and beyond during the Middle Bronze Age (ap- riod that witnessed the blossoming of Middle Bronze Age city proximately 2000 to 1550 BCE), Late Bronze Age (approxi- states, periods of Egyptian domination and retreat during the Late mately 1550 to 1200/1150 BCE), and Early Iron Age/Iron Age I Bronze Age, and the emergence of the so-called at the (approximately 1200/1150 to 1000 BCE). Here we aim to explore onset of the Early Iron Age during the 12th century BCE the transformation of eastern Mediterranean cuisine as a con- sequence of Bronze Age globalization by analyzing microscopic Results and molecular traces of food remains in human dental calculus, a Plant remains were abundant in the Megiddo and Tel Erani calcified form of dental plaque, from the Southern Levant during dental calculus, and plant microremains and proteins were - the second millennium BCE (Table 1). We focus on two sites, served in all 16 dental calculus samples. Of these, probable di- the Middle to Late Bronze Age urban center of Megiddo (17th etary microremains were identified in all 16 analyzed samples (SI to 15th centuries BCE) and the Early Iron Age site of Tel Erani Appendix, Table S1), and included phytoliths consistent with (11th century BCE) (Fig. 1). In total, we analyze dental calculus wheat (Triticum), panicoid/millet (Panicoideae), and date palm from 16 individuals: 13 from Megiddo (SI Appendix, Fig. S2) and (Phoenix sp.). Dietary proteins were observed in 5 of 14 analyzed 3 from Tel Erani (SI Appendix, Fig. S3). specimens (SI Appendix, Table S2), and consisted of 19 dietary

Scott et al. PNAS | 3of10 Exotic foods reveal contact between South Asia and the Near East during the second mil- https://doi.org/10.1073/pnas.2014956117 lennium BCE Downloaded by guest on October 1, 2021 Table 1. Overview of dietary findings Individual Burial Context* Microremains Dietary proteins

Megiddo, Middle Bronze Age III to Late Bronze Age I, n = 13 MGD001 Tomb 50 “King”; elite masonry Poaceae, cf. Triticum, Phoenix (date palm), None (triple burial); ca. 1650–1550 BCE Arecaceae unspecific (palm), bark, eudicot MGD002 Tomb 50 “Queen”; elite masonry chamber tomb Poaceae, eudicot None (triple burial); ca. 1650–1550 BCE MGD006 Tomb 16/H/45 Double pit burial; ca. 1550–1450 BCE Poaceae, eudicot None MGD007 Tomb 16/H/45 Double pit burial; ca. 1550–1450 BCE Poaceae None MGD008 Tomb 12/K/89 Pit burial; 1496–1320 BCE Poaceae, eudicot None MGD009 Tomb 12/K/96 Double pit burial; ca. 1650–1400 BCE Poaceae, cf. Triticum, Triticeae, eudicot None MGD010 Tomb 12/K/96 Double pit burial; 1638–1413 BCE Poaceae, Triticeae, eudicot None MGD011 Tomb 14/K/ Stone-lined , exotic ; Poaceae, cf. Triticum, Phoenix (date palm), Sesamum, 119, lower 1688–1535 BCE eudicot Triticum/Aegilops MGD013 Tomb 10/K/118 Triple pithos burial; ca. 1650–1400 BCE Poaceae, Phoenix (date palm), eudicot, bark Not analyzed MGD016 Tomb 14/K/49 Double brick-lined pit Burial; 1509–1432 BCE Poaceae, eudicot None MGD017 Tomb 100 Masonry chamber tomb with corbelled roof, Poaceae, Cyperaceae, Phoenix (date palm), None many commingled individuals; exotic grave Arecaceae (palm), eudicot goods; ca. 1650–1400 BCE MGD018 Tomb 100 Masonry chamber tomb with corbelled roof, Poaceae, eudicot, Triticeae Curcuma, Glycine many commingled individuals; exotic grave goods; ca. 1630–1550 BCE MGD021 Tomb 50 Elite masonry chamber tomb (triple burial); ca. Poaceae, Triticeae, Phoenix (date palm), Not analyzed 1650–1550 BCE eudicot Tel Erani, Early Iron Age, n = 3 ERA005 Burial L2091 Burial offering of one juglet; ca. 1100–1000 BCE Poaceae Sesamum ERA017 Burial L2160 Burial offering of one Flask; ca. 1100–1000 BCE Poaceae, Cyperaceae, eudicot, panicoid cf. Musa millet ERA023 Burial L2181 No burial offerings; ca. 1100–1000 BCE Poaceae, eudicot Sesamum

*Dates are provided as relative dates (marked with “ca.”) or calibrated radiocarbon dates (2σ). See Dataset S1.

proteins from cereals (Triticum/Aegilops), oilseeds (Sesamum, or gymnosperm), were identified in 13 individuals (SI Appendix, Glycine), fruits (Musa), and spices (Curcuma). Fig. S7), totaling 245 phytoliths (SI Appendix, Table S1). Starch granules were rare in all samples, except ERA23 (SI Appendix, Microremains. Microremains were analyzed from all 16 individ- Fig. S4), even in samples not exposed to heat during protein uals, including 2 individuals (MGD013 and MGD021) with in- extraction, and few could be identified. However, individual sufficient calculus to conduct proteomic analysis (SI Appendix, starch granules consistent with Triticeae were found in ERA023, Table S1 and Datasets S2 and S3). Representative examples of MGD018, and MGD021 (SI Appendix, Table S1). In addition, we the dietary microremains observed in the Megiddo and Tel Erani found 26 other types of microremains whose origins are either dental calculus are provided in Fig. 3. Both the Megiddo and Tel ambiguous (pollen, fungal particles, diatoms, foraminifera, sponge Erani individuals produced microremains assemblages consisting spicules, , fibers, insect fragments) or possible contami- of three main components: 1) Dietary microremains, most no- nants (skin scales) (Datasets S2 and S3). These microremains were tably phytoliths (84 total morphotypes, 4,983 phytoliths), but also not further analyzed. starches (333 granules); 2) nondietary remains, such as fibers; Among phytoliths assigned to Poaceae, all morphotypes that and 3) ambiguous microremains, such as fungal particles, char- form in grasses were represented, including those forming in particles, and other remains of unknown or ambiguous or- epidermal short cells, long cells, bulliforms, hairs, papilla, and igin (Dataset S2). The abundance of microremains observed stoma (Dataset S2), in both single-cell and articulated forms. across individuals was highly variable (SI Appendix, Fig. S4), Grass phytoliths predominantly belonged to the pooid group ranging from 9 (MGD007) to 1,795 (MGD001). (n = 432), but a smaller number of single-cell short cells could be Phytoliths, a robust and abundant type of plant microfossil, assigned to the chloridoid (n = 51) and panicoid (n = 10) grass allow the identification of vascular plants, and among angio- types (SI Appendix,Fig.S8). Although most phytoliths could not be sperms (flowering seed plants), they primarily form in monocots, assigned to specific wild or domesticated sources, the assemblage is particularly in members of the plant family Poaceae (grasses, consistent with grain consumption, and 105 articulated dendritic types including cereals). In contrast, magnoliids, eudicots, and other were identified in 8 individuals (ERA017, MGD001, MGD002, angiosperms (which include most fruits and vegetables) tend to MGD009, MGD010 and MGD011, MGD017, and MGD021). form few phytoliths, while gymnosperms (nonflowering seed Among these, 10 articulated husk phytoliths consistent with plants) produce even fewer (67). The dental calculus phytoliths Triticum (wheat) were identified in MGD001, MGD009, and observed in this study corresponded to these expected propor- MGD011, and 5 short cells (wide lobed bilobates) deriving from tions (SI Appendix, Figs. S4 and S5) and were predominantly Panicoideae grasses (e.g., ) were found in ERA017 (SI derived from the leaves, stems, and husks of wild or domestic Appendix, Table S1 and Dataset S2). Poaceae (SI Appendix, Fig. S6), totaling 2,551 phytoliths, plus an Cones representing sedge (Cyperaceae) leaf were found in additional 916 unspecific Poaceae phytoliths (SI Appendix, Table MGD001, MGD017, MGD021, and ERA017 (SI Appendix, Fig. S1). Smaller numbers of morphotypes from other monocots S7 and Table S1). Such cones form in a variety of sedge cells, in- such as Cyperaceae (sedges) and Arecaceae (palms), as well as cluding sedge achenes and achene bracts cells. Palm (Arecaceae) eudicot fruit/leaf phytoliths and nondiagnostic bark (angiosperm phytoliths were identified in five individuals (SI Appendix,TableS1).

4of10 | PNAS Scott et al. https://doi.org/10.1073/pnas.2014956117 Exotic foods reveal contact between South Asia and the Near East during the second millennium BCE Downloaded by guest on October 1, 2021 A total of 12 globular rugulate/echinate phytoliths of date palm ∼80% of the total proteins in whole wheat flour, and LMW (Phoenix) were observed in MGD001, MGD011, MGD013, glutenin accounts for 20 to 35% of the gluten protein content (71). MGD017, and MGD021, and phytoliths from another unknown Sesame proteins were identified in one individual from species of palm were also detected in MGD001 and MGD017 Megiddo (MGD011) and two from Tel Erani (ERA005, ERA023) (Datasets S1 and S2). Globular echinate palm leaf phytoliths are and consisted of 2 proteins supported by a total of 29 peptides and known to be a particularly resilient morphotype (68). Finally, 217 78 PSMs: 11S globulin and 2S albumin seed storage proteins. plate, jigsaw, sclereid, tracheid, and related phytoliths from These two seed storage proteins are tissue-specific and expressed eudicot epidermal tissue were identified in 14 individuals, in- during seed maturation. Together, they make up most of the cluding a single Megiddo individual (MGD001) with 3 jigsaw protein in sesame seeds, accounting for 60 to 70% and 15 to 25% morphotypes displaying protuberance decorations thought to of total seed protein, respectively (72). originate from the eudicot epidermal tissue of fruit and seeds (SI Soybean was identified in a single Megiddo individual (MGD018) Appendix, Fig. S7 and Datasets S2 and S3) (69). and is supported by 30 PSMs specific to the proteins glycinin, an 11S storage protein, and β-conglycinin, a 7S storage protein. These Proteomics. Total protein was extracted from the dental calculus two proteins are the primary storage proteins in soybean seeds, of 14 individuals. Protein recovery was variable, but all samples and together they make up ∼65% of the total soybean protein yielded proteins typical of an oral microbiome (SI Appendix, Fig. content (73). In addition to these two seed storage proteins, we S9 and Dataset S4) and contained damage-associated modifi- also identified two peptides supported by two PSMs specific to cations (N,Q deamidation) (SI Appendix, Table S3) consistent soybean sucrose-binding protein, another member of the seed with ancient samples. Dietary proteins were identified in 5 in- dividuals and consisted of 19 proteins from 5 plants of known storage protein superfamily. We identified endo-1,3-β-glucanase, an enzyme important in dietary importance: wheat (Triticum/Aegilops), sesame (Ses- amum), soybean (Glycine), banana (probable Musa), and tur- fruit ripening, in a single individual from Tel Erani (ERA017). meric (Curcuma) (Fig. 4 and SI Appendix, Fig. S10 and Table This protein was supported by two peptides, of which one is S2). Detailed information regarding the identified peptides and highly specific to banana (Musa or Ensete), while the second is peptide spectral matches (PSMs) for each protein is provided in found in Musa and a number of other flowering plants, but not Dataset S5. Ensete. Of the two plants, only members of Musa produce edible Wheat proteins were identified in a single individual from fruits, while Ensete (Abyssinian banana) is consumed for its Megiddo (MGD011) and consisted of two major seed proteins: starchy pseudostem and corm. The fruit of ripe bananas contains α-amylase inhibitor (AAI) and low molecular weight (LMW) few proteins but relatively high concentrations of β-glucanases ANTHROPOLOGY glutenin. AAI makes up ∼4% of the protein content of wheat (74), which are highly resistant to heat and proteolytic degra- and is highly resistant to heat or proteolytic digestion (70). The dation (75). Because this protein is expressed in fruit and peel identification of AAI was supported by three PSMs, with two and increases in abundance as the fruit ripens (76, 77), we ten- PSMs matching specifically to either Triticum aestivum or Aegi- tatively identify this enzyme as originating from Musa. However, lops tauschii (a wild progenitor of T. aestivum), while the third the enzyme is also expressed in generalized stress response, and peptide was less specific and is also found in Hordeum (barley). thus is also present in diseased and injured plant tissues. Taken LMW glutenin, a major gluten protein, was identified with the together, the identification of Musa is most strongly supported, support of 3 peptides and 13 PSMs. Gluten proteins make up but Ensete cannot be fully excluded.

Fig. 3. Microremains in Megiddo and Tel Erani dental calculus. (A) Articulated Poaceae husk phytolith, identified as wheat (MGD001). (B) Poaceae stem/leaf phytolith (MGD001). (C) Poaceae short cell rondel (MGD001). (D) Wide-lobed bilobate short cell identified as panicoid (ERA017). (E) Poaceae polylobate short cell (MGD001). (F) Cone phytolith identified as sedge leaf (MGD018). (G) Spheroid echinate identified as date palm (MGD001). (H) Spheroid echinate phy- tolith, identified as nondiagnostic palm (MGD001). (I) Polyhedral plate phytolith, identified as eudicot (MGD011). (J) Decorated jigsaw phytolith, likely from fruit (MGD001). (K) Spheroid psilate phytolith, identified as bark type (MGD001). (L) Damaged Triticeae starch in brightfield, with Inset showing an absence of birefringence in cross-polarized light (MGD010). (Scale bars, 20 μm.)

Scott et al. PNAS | 5of10 Exotic foods reveal contact between South Asia and the Near East during the second mil- https://doi.org/10.1073/pnas.2014956117 lennium BCE Downloaded by guest on October 1, 2021 Finally, we identified the turmeric protein turmerin, supported which they derive remain untranslated or unknown; for example, by three peptides and four PSMs, in a single individual from it is still disputed whether Egyptian “ baq oil” refers to oil, Megiddo (MGD018). Turmerin is an α-amylase/trypsin inhibitor moringa oil, or to the oil of another plant (103). There was high that belongs to the leguminous kunitz-type serine inhibitor family of proteases. It has known antioxidant activity (78) and also plays a role in plant defense. Although making up only 0.1% of the dry-weight of turmeric, it is one of the most stable tur- A meric proteins, being resistant to heat, digestive enzymes, and UV irradiation (78). Discussion Among the identified dietary taxa, wheat and date palm were expected finds, as wheat has been a staple crop in the Levant since the seventh millennium BCE (79) and date palm fruits have been consumed and traded since at least the fifth millennium BCE (54, 57, 80). Moreover, date palm seeds and leaves have been previously recovered from Late Bronze II burials at Megiddo (81). Microfossil remains of both crops have been previously identified in the Bronze Age Levant (82–84), and here B we identified both wheat and date palm phytoliths, as well as wheat proteins, including a major wheat gluten. Sesame and millet, although not unexpected, are important new finds. These nonlocal domesticates from South and East Asia, respectively, spread to West Asia during the Bronze Age (85), but their arrival in the Levant is less well understood. To date, the oldest remains of sesame seeds (Sesamum indicum) have been found at Harappan sites in the Indus Valley (2500 to 2000 BCE), but charred and desiccated seeds have also been sporadically recovered at sites in the Near East since the late third millennium BCE (86, 87). The Akkadian word “šamaš- šammȗ,” which refers to an oil plant (possibly sesame), appears C in cuneiform texts from 2400 BCE onwards (87–89), and al- though questioned in the past, the identification of sesame seeds in the tomb of Tutankhamun (14th century BCE) is now con- sidered credible (90). We found robust evidence for multiple Sesamum proteins in individuals at both Megiddo and Tel Erani, suggesting that by the second millennium BCE, sesame had be- come a staple oil-bearing crop in the Levant. We identified Panicoideae phytoliths consistent with dietary millets in a single individual at Tel Erani. Although not identi- fiable below the taxonomic level of subfamily, the Eurasian grasses Setaria and Panicum and the African grasses Sorghum D and Pennisetum are possible candidates. Among these, Sorghum and Pennisetum are unlikely, as there is no evidence for the dispersal of sorghum into the Levant prior to the medieval pe- riod (54), and although African pearl millet (Pennisetum glau- cum) had spread from to South Asia by the mid- second millennium BCE (91, 92), there is no evidence for its cultivation in the Near East. In contrast, Asian broomcorn (Panicum miliaceum) and foxtail (Setaria italica) millets are known to have reached western Eurasia via (93, 94) by the second or possibly third millennium BCE (54, 95), al- though likely no earlier based on recent and collagen stable studies (96). Within the Near East, cur- E rent macrobotanical evidence suggests that S. italica and P. miliaceum millets functioned as minor crops from the first mil- lennium onwards (54, 97–100), and this is consistent with our identification of the panicoid phytoliths at Iron Age Tel Erani. In contrast to the crops above, the consumption of soybean (Glycine), probable banana (Musa), and turmeric (Curcuma longa) were unexpected finds. Soybean cultivation was unknown in this region before the CE and, like millet, its domestication center was near the in Central China, where it was cultivated as early as 7000 to 6500 BCE (101). However, soybean, like sesame, is a major oil plant, and its Fig. 4. Representative MS/MS spectra of selected dietary peptides. (A) oil could have been transported over long distances. Exotic oils Sesamum, 11S globulin protein (MGD011). (B) Triticum/Aegilops, LMW glu- are frequently mentioned in Old Babylonian, Akkadian, and tenin (MGD011). (C) Musa, β-1,3 glucanase (ERA017). (D) Curcuma, turmerin Egyptian texts (102–104). However, many of the plants from (MGD018). (E) Glycine, sucrose-binding protein (MGD018).

6of10 | PNAS Scott et al. https://doi.org/10.1073/pnas.2014956117 Exotic foods reveal contact between South Asia and the Near East during the second millennium BCE Downloaded by guest on October 1, 2021 demand for oils of different flavor and origin in ancient Meso- The Asian tropical plant family Zingiberaceae contains numerous potamia and Egypt, where such oils played vital roles in cuisine, economically useful plants used for food, spices, , dyes, medication, bodycare, and illumination. Furthermore, they were and perfumes (122, 123). Among these, the rhizomes of the genus part of a broad spectrum of daily and ritual practices, where Curcuma are consumed as foods in South and , they were used, for example, to anoint objects and embalm the of which domesticated turmeric (C. longa) is among the most dead (103–106). Recent X-ray computed tomography imaging important and widely used. Turmeric starch grains have been of archaeological soybeans indicates that oil content was a identified in both cattle dental calculus and at the major target of selection during domestication, and cultivars Harappan site of Farmana, dating to between 2600 and 2200 with high oil content were prevalent in China by approximately BCE (124–126). Turmeric has multiple uses as both a spice and 2000 BCE (107). The relative scarcity of evidence for soybean a cloth dye (72), and early medical texts in China and India also in the archaeological record might be explained by its pre- report its use as a medicinal (127). Within the Near East, the dominant use as an oil combined with a lack of organic residue earliest references to turmeric appear during the seventh cen- analysis on relevant material and/or the difficulty of differen- tury BCE in Assyrian cuneiform medical texts from Ashurba- tiating plant oils through lipid analysis. Proteomics has been nipal’s library at Nineveh (128), but no archaeological evidence previously shown to be a powerful for identifying archae- has been found prior to the Islamic period during the 11th to ological oils and fats (108), as plant oils and rendered animal 13th centuries CE (1). Our identification of Curcuma protein at fats produced using preindustrial techniques generally contain the site of Megiddo suggests that it was already present or ac- residual proteins. By identifying strong evidence of soybean cessible to individuals in the Levant as early as the mid-second seed proteins in dental calculus, we confirm that Megiddo indi- millennium BCE. Interestingly, turmeric protein was identified vidual MGD018, who was also buried with other exotic grave goods at Megiddo in the same individual (MGD018) whose dental (Table 1 and SI Appendix), likely had access to soybean oil, and we calculus contained soybean protein. This individual was buried further demonstrate the utility of proteomics as a method for in a wealthy collective tomb containing exotic goods (Table 1 identifying exotic oils in the Near East that otherwise leave few and SI Appendix), suggesting that this individual was either well archaeological traces. connected with trading activities or may have even been a Banana (Musa) was domesticated during the fifth millennium merchant or trader themself. As such, the individual may have BCE in New Guinea (109) and by the first millennium BCE had consumed foods seasoned with turmeric or prepared with soy dispersed under human cultivation as far west as Cameroon in oil in the Levant, in South Asia, or elsewhere. West Africa (110, 111). However, reconstructing the intervening Overall, the plant microremains and proteins identified in cultivation and trade of bananas has proven particularly difficult dental calculus from Megiddo and Tel Erani point toward the ANTHROPOLOGY to trace through the archaeological record. Banana fruit is highly existence of a dynamic and complex exchange network connecting perishable and domesticated bananas are seedless (reproducing the Mediterranean with South Asia during the second millennium instead by cuttings), and thus there is a strong bias against the BCE that outlived the dramatic socio-political transformation and recovery of banana macrobotanical remains. Phytoliths represent associated shift from centrally organized trade during the Bronze the principle method employed to trace early banana use, but Age to diverse small-scale trade entrepreneurship from the Early banana phytolith recovery from archaeological contexts is gen- Iron Age onwards. Historically, archaeologists and historians in the erally low, perhaps due to the limited number of phytoliths Near East have relied on texts, iconography, and macrobotanical produced by the plant, the lack of phytoliths in the consumed remains as their primary sources of information in reconstructing mesocarp, and other taphonomic factors (112, 113). Direct evi- the region’s cuisine and trade connections. However, these ap- dence of banana, in the form of phytoliths, has been found at proaches alone can be limiting. Although the text base is rich, three Indus sites dating to the late third millennium (114–117), including both cuneiform texts and papyri, many of the attested placing the crop in South Asia by that date. The earliest plant-related terms cannot yet be translated or fully understood, reported archaeobotanical evidence for banana in the Near despite being transliterated. Iconography, while providing vivid East is desiccated fruit pulp recovered from a vessel in an images of the past, can lack the botanical detail necessary to make Egyptian 18th Dynasty tomb (15th ct. BCE), but the identifi- conclusive identifications, and macrobotanical remains are gener- cation is highly contested (118, 119). Other scholars have ar- ally strongly biased toward grains and other seeds, especially those guedforanintroductionofbananaintheArabianpeninsula that have been carbonized. While important, these methods can prior to the ninth century BCE (120), but the first secure miss oils, fruits, and spices that are unlikely to carbonize and which identification of Musa in the Near East consists of a banana leaf may have been traded in small quantities or in already processed found in an Egyptian tomb at Antinoë dating to the fifth cen- powdered or liquid forms. tury CE (121), long after the crop had already spread further Here we demonstrate the utility of combining microscopic and westtobecomeastapleinWestAfrica(110,111).Thepre- molecular techniques to reveal a broader range of foodstuffs that historic spread and use of banana in South and West Asia re- include both staple grains, as well as oils, fruits, and spices that mains poorly known, in large part due to preservation biases. otherwise leave behind little macrobotanical evidence. In par- Our identification of a major banana fruit-ripening protein in ticular, we found that plant phytoliths primarily reflected bulk the dental calculus of individual ERA017 at Tel Erani lends dietary items, and were dominated by high levels of grass phy- support for either the banana being present in the Levant by the toliths, most likely deriving from wheat, but also including first millennium BCE or a mobile individual (e.g., a merchant or probable millets. We also identified palm phytoliths, almost seafarer) who consumed banana during his lifetime in South or certainly from date fruit. All of these resources are monocots, East Asia before being buried at Tel Erani. This identification, reflecting their tendency to be well represented by phytoliths. In although only supported by two peptides, is nevertheless specific, contrast, protein analysis revealed a wider diversity of foods and and within the broader banana family Musaceae the peptides are is better at identifying plants with low levels of silicification and consistent only with Asian bananas (Musa), to the exclusion of therefore few phytoliths, especially if they are protein-rich. Al- other related plants, such as the African enset (Ensete), also though the modes of protein preservation and incorporation in known as the Abyssinian banana. The protein itself, which is ancient dental calculus are still under study, most of the proteins abundant in ripening fruits, is also supportive of a Musa fruit we identified were either protease inhibitors or belonged to the identification, as the fruits of the Abyssinian banana are inedible seed storage protein superfamily. Both types of proteins are and only the starchy portions of the pseudostem and corm highly stable against proteolysis and thermal processing, traits are consumed. that may increase their likelihood of survival in archaeological

Scott et al. PNAS | 7of10 Exotic foods reveal contact between South Asia and the Near East during the second mil- https://doi.org/10.1073/pnas.2014956117 lennium BCE Downloaded by guest on October 1, 2021 calculus. These same traits can also contribute to their potential Primatology at the Max Planck Institute for Evolutionary Anthropology, allergenicity (129), and many of the identified proteins are also Leipzig. Samples were left in EDTA until decalcification was complete, which known allergens. In addition to being resilient, several of the varied from a few hours to a few days. In two samples (MGD002 and proteins we identified, especially seed proteins, are also highly MGD016), we performed a predecalcification to separate microremains on the outside of the calculus pieces from the interior (staged decalcification) abundant in the foods from which they originate, and similar (SI Appendix). The samples were then centrifuged at 2,000 × g for 10 min seed storage proteins have been previously identified in Neo- (Roth Minicentrifuge) and EDTA was removed from the samples by pipetting lithic pottery residues (130) and in dental calculus from the the supernatant. This process was repeated three times. Then 25% glycerine medieval and postmedieval periods (131–133). for mounting was added to the tube. In addition to analyzing whole calculus The identification of sesame, soybean, banana, and turmeric pieces, cellular debris pellets left over after protein extraction (see next sec- proteins in Megiddo and Tel Erani dental calculus points to the tion) were also analyzed for ERA005, ERA017, ERA023, MGD001, MGD002, μ need to reevaluate the current evidence for the second millen- MGD009, MGD010, MGD011, and MGD017. For these pellets, 100 Lof25% nium BCE Indo-Mediterranean trade. Previous suggestions of glycerine solution was slowly added to the tubes to avoid spillage loss due to foaming of residual SDS from protein extraction. For all samples, 20 μLofeach such a trade network between India and Egypt were largely ig- sample were mounted on slides with 18 × 18- or 22 × 22-mm coverslips nored due to the limited physical evidence (119, 134). However, depending on volume. The mounting was performed in a laminar flow hood such evidence is now growing, and thus earlier identifications of and examined under brightfield and cross-polarized light on a Zeiss Axioscope plants (e.g., jasmine, nutmeg, cinnamon), both from archaeologi- microscope at 400× magnification (Num. Aperture = 0.95). Microremains were cal remains and from texts, should be reconsidered. The broader analyzed by examining the whole slide and any encountered microremains body of evidence for exotic goods, which also includes zebu cattle, were photographed, described, and documented using the procedures de- chickens, citron, melon, cloves, millet, vanillin, peppercorns, scribed in SI Appendix, Evidence of Long-Distance Trade in the Ancient World, monkeys, and beetles, points to a pattern of established trade. The and Procedures for Microremains Analysis. individuals from Megiddo, tomb 100 (MGD017, MGD018) not Proteomics. Protein extractions were performed on 14 dental calculus samples only had access to exotic food, but were also buried with precious using a filter-aided sample preparation protocol modified for ancient pro- grave goods (Dataset S2), which might indicate a higher-status teins (see published protocol at https://www.protocols.io/view/ancient- position or the collective burial of members of a trading house. proteins-extraction-protocol-7vwhn7e). Samples were extracted and diges- Such traders and travelers may have transported cargoes of ani- ted alongside negative extraction blanks in order to monitor for potential mals, spices, dried fruits, oils, and perfumes via different routes, laboratory contamination. Cellular debris pellets left over after protein ex- either overland through Iran and the Central Asian oases or by sea traction were set aside for microscopic examination of microremains (see across the Indian Ocean to either the Red Sea or Persian Gulf, above section). Extracted peptides were analyzed by LC-MS/MS using a or both. Q-Exactive mass spectrometer (Thermo Scientific) coupled to an ACQUITY UPLC M-Class system (Waters) at the Functional Genomics Center Zurich of This study highlights the potential of microscopic and molecular the University/Eidgenössiche Technische Hochschule Zurich (SI Appendix). methods to reveal elements of trade and cuisine that otherwise Injection blanks were also run between each sample in order to identify and leave few archaeological traces. As detection methods continue to reduce potential carryover across samples. MS/MS spectra were converted to improve, there will likely need to be a fundamental reconsidera- Mascot generic files by MSConvert v3.0.11781 using the 100 most-intense tion of the dimensions and complexities of Bronze Age trans- peaks in each spectra. All MS/MS samples were analyzed using Mascot (Matrix Eurasian trade. Although named for a that is highly visible Science; v2.6.0). Mascot was set up to search the SwissProt Release 2019_08 in the archaeological record, the process of Bronzization was likely database (560,823 entries) and Uniprot Trembl 2017_07 (88,032,926 se- a much broader phenomenon that also linked cuisines and econ- quences) assuming the digestion enzyme trypsin, with an automatic decoy option. Mascot was searched with a fragment ion mass tolerance of 0.050 Da omies across Eurasia. and a parent ion tolerance of 10.0 PPM. Carbamidomethyl of cysteine was specified in Mascot as a fixed modification. Deamidation of asparagine and Materials and Methods glutamine and oxidation of methionine and proline were specified in Mascot Excavation. All individuals analyzed in this study were excavated and docu- as variable modifications. All protein identifications were established at a mented in their archaeological context from the sites of Megiddo and Tel protein false-discovery rate of less than 3.0% and peptide false-discovery rate Erani within the last decade (Dataset S1). An archaeological and anthropo- of less than 1.0% using the Protein Prophet algorithm (137) implemented in logical overview of each burial is provided in SI Appendix. Burials have been the software program Scaffold (version Scaffold_4.8.9; Proteome Software). dated using radiometric methods and/or associated grave goods. Dietary proteins were filtered at a minimum of 95% protein identification probability. Additionally, only proteins with a minimum of two unique Sampling. Dental calculus sampling was performed in a clean laboratory at peptides with at least one species-specific peptide were accepted (Dataset the Tel Aviv University Megiddo excavation archives and at the Tel Erani S5). Proteins that contained similar peptides that could not be differenti- excavation storage facility of the Israel Antiquity Authorities. Nitrile gloves ated on the basis of MS/MS analysis alone were grouped to satisfy the were worn during collection, and calculus was sampled using dental curettes principles of parsimony. Proteins sharing significant peptide evidence that were replaced or cleaned with isopropanol between samples. Calculus were grouped into clusters. No dietary proteins were found in the ex- was collected onto weighing paper or aluminum foil and packaged indi- traction blanks, which contained only reagents, such as porcine trypsin, vidually. Samples were received at the Max Planck Institute for the Science of and known laboratory contaminants, such as collagen, keratin, or bacterial ancient proteomics laboratory, where they were weighed and proteins. Representative MS/MS spectra of selected dietary proteins are subsampled prior to protein extraction. Approximately 5 to 10 mg of dental provided in SI Appendix,Fig.S10. A complete list of all identified dietary calculus was used for protein analysis, and 0.5 to 2 mg of calculus was spectra is provided in Dataset S5. subsampled and transferred into 1.5-mg Eppendorf tubes for analysis by microscopy. These samples, along with nine pellets remaining after protein Data Availability. Protein spectra have been deposited to the ProteomeXchange extraction were transported to the Max Planck Institute for Evolutionary Consortium via the PRIDE partner repository (https://www.ebi.ac.uk/pride) Anthropology for analysis of microremains. under the dataset identifier PXD021498.

Microremains. Prior to analysis, we first tested for the presence of surface ACKNOWLEDGMENTS. This study was funded by the European Research contaminants in a subset of samples (n = 2; MGD002, MGD016) by per- Council (ERC) under the European Union’s Horizon 2020 research innovation forming a staged decalcification to compare the relative abundance and programme (ERC-2015-StG 678901-Food-Transforms) as part of P.W.S.’sERC composition of surface and interior microremains (135, 136). We found Starting Grant project “FoodTransforms: Transformations of Food in the East- ” surface contamination to be negligible (SI Appendix, Fig. S11), and so pro- ern Mediterranean Late Bronze Age. Work at Megiddo is currently supported by the Shmunis Family Foundation, the Dan David Foundation, Jacques Chahine, ceeded to process the remaining samples. Mark Weissman, as well as Norman and Vivian Belmonte. We thank Zeev Herzog ∼ To extract the microremains we added 1 mL of 0.5 M EDTA to decalcify and Lily Avitz Singer for giving us the permission to publish the pottery our preweighed dental calculus samples that were in 1.5-mL Eppendorf sherd photograph and for supplying information as to its context; and Chen tubes under a Bio Air Aura Mini laminar flow in the Department of Tao and Robert Spengler for helpful comments on the manuscript.

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